1. Chapter 32

2. Defining Fungi Mycologists believe there may be as many as 1.5 million fungal species Single-celled or multicellular Sexual or asexual Specialized to extract and absorb nutrients from surroundings Animal and fungi last shared a common ancestor 460 MYA Some debate on timing May have 670 MYA ancestor 2

3. 7 monophyletic phyla Microsporidia Blastocladiomycota Neocallismastigamycota Chytridiomycota Glomeromycota Basidiomycota Ascomycota 1 polyphyletic Zygomycota 3

4. 4 Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. c. MicrosporidiaBlastocladiomycotaZygomycotaNeocallimastigomycotaChytridiomycotaGlomeromycotaBasidiomycotaAscomycota Dikarya Fungi a. 10 μm b.h.g. 300 μm200 μm300 μm e.f.d. 300 μm500 μm a: © Dr. Ronny Larsson; b: Contributed by Don Barr, Mycological Society of America; c: © Carolina Biological Supply Company/Phototake; d: Contributed by Don Barr, Mycological Society of America; e: © Dr. Yuuji Tsukii; f: © Yolande Dalpe, Agriculture and Agri-Food Canada; g: © inga spence/Alamy; h: © Michael & Patricia Fogden

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6. General Biology of the Fungi Multicellular fungi consist of long, slender filaments called hyphae Some hyphae are continuous Others are divided by septa Cytoplasm flows throughout hyphae Allows rapid growth under good conditions 6

7. 7 Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Hypha Nuclei Pore Septum 0.2 µm Dikaryotic cell Septa with pores (right): © Garry T. Cole/ Biological Photo Service

8. Mycelium – mass of connected hyphae Grows through and digests its substrate Fungal cell walls include chitin Also found in the hard shells (exoskeletons) of arthropods 8 Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Hyphae Mycelium 10 μm (inset): © Micro Discovery/Corbis; (right): © Michael & Patricia Fogden/Corbis

9. Hyphae may have more than one nucleus Monokaryotic – 1 nucleus Dikaryotic – 2 nuclei Both genomes transcribed Sometimes many nuclei intermingle in the common cytoplasm of the fungal mycelium Heterokaryotic – nuclei from genetically distinct individuals Homokaryotic – nuclei are genetically similar to one another 9

10. Fungi have an unusual mitosis Cell is not relevant unit of reproduction Nuclear envelope does not break down and re-form Instead, the spindle apparatus is formed within it Fungi lack centrioles Spindle plaques regulate microtubule formation during mitosis 10

11. Reproduction Capable of both sexual and asexual reproduction Sexual reproduction Fusion of two haploid hyphae of compatible mating types In some fungi, fusion immediately results in a diploid (2n) cell Others, have a dikaryon stage (1n + 1n) before parental nuclei form diploid nucleus May form mushrooms or puffballs 11

12. Spores are the most common means of reproduction among fungi May form from sexual or asexual processes Most are dispersed by wind 12 Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. 10 μm © Eye of Science/Photo Researchers, Inc.

13. Nutrition Obtain food by secreting digestive enzymes into surroundings Then absorb the organic molecules produced by this external digestion Great surface area-to-volume ratio Fungi can break down cellulose and lignin Decompose wood Some fungi are carnivorous 13 b. Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. © L. West/ Photo Researchers, Inc. Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. a. 400 µm Fungus Nematode Fungal loop © Carolina Biological Supply Company/Phototake

14. Microsporidia Obligate, intracellular, animal parasites Long thought to be protists Lack mitochondria Ancestors lost them 14 Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Neocallimastigomycota Zygomycota Chytridiomycota GlomeromycotaBasidiomycota Ascomycota Blastocladiomycota Microsporidia Fungi

15. Encephalitozoon cuniculi Commonly cause disease in immunosuppressed patients Infect hosts with their spores, which contain a polar tube Infects intestinal and neuronal cells, leading to diarrhea and neurodegenerative disease 15 Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. 0.5 μm Spore Polar tube (left): © Daniel P. Fedorko

16. Chytridiomycota Chytridiomycetes or chytrids Aquatic, flagellated fungi Closely related to ancestral fungi Have motile zoospores Batrachochytrium dendrobatidis has been implicated in amphibian die-offs 16 NeocallimastigomycotaZygomycotaChytridiomycota GlomeromycotaBasidiomycota Ascomycota Blastocladiomycota Microsporidia Fungi Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

17. 17 Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Zoospore Contributed by Daniel Wubah, Mycological Society of America Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Chytrid Oedogonium filament Contributed by Don Barr, Mycological Society of America

18. Blastocladiomycetes Uniflagellated zoospores Allomyces example Water mold Haplodiplontic life cycle Female gametes secrete pheromone to attract male gametes Giant mitochondria in its zoospores 18

19. 19 Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. 100 µm Female gametangium Male gametangium Haploid gametes (n) Sporophyte(2n) Gametophyte (n) Mature sporophyte (2n) Asexual sporangium Sporophyte Sporangium (2n) n 2n2n a.b. Young gametophyte (n) Haploid zoospore (n) Young sporophyte (2n) Young sporophyte (2n) Asexual reproduction Diploid zoospores (2n) Mature gametophyte (n) FERTILIZATION MEIOSIS MITOSIS a: © Carolina Biological Supply Company/Phototake

20. Neocallimastigomycota Digest plant biomass in mammalian herbivore rumens Mammal depends on fungi for sufficient calories Greatly reduced mitochondria lack cristae Zoospores have multiple flagella Horizontal gene transfer brought cellulase gene from bacteria into Neocallimastix genome 20

21. Zygomycota Zygomycetes are incredibly diverse Not monophyletic – still under research Include the common bread molds A few human pathogens 21 Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Neocallimastigomycota ZygomycotaChytridiomycota GlomeromycotaBasidiomycota Ascomycota BlastocladiomycotaMicrosporidia Fungi

22. Sexual reproduction Fusion of gametangia Haploid nuclei fuse to form diploid zygote nuclei – karyogamy Develops into zygosporangium in which zygospore develops Meiosis occurs during germination of zygospore Releases haploid spores Asexual reproduction more common Sporangiophores have sporangia that release spores 22

23. 23 a. b. Rhizoid Sporangiophore Sporangium – Mating strain n n + n (+)(–) Zygosporangium Hypha Gametangia Spores MEIOSIS 2n 700 µm Asexual reproduction (Meiosis occurs during germination) FERTILIZATION KARYOGAMY GERMINATION Germinating zygosporangium + Mating strain GERMINATION Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. a: © Carolina Biological Supply Company/Phototake

24. Glomeromycota Glomeromycetes are a tiny group of fungi Form intracellular associations with plant roots called arbuscular mycorrhizae Cannot survive in absence of host plant No evidence of sexual reproduction 24 Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Neocallimastigomycota Zygomycota ChytridiomycotaGlomeromycotaBasidiomycota Ascomycota Blastocladiomycota Microsporidia Fungi

25. Basidiomycota Basidiomycetes are some of the most familiar fungi Mushrooms, toadstools, puffballs, shelf fungi, etc. Also important plant pathogens like rusts and smuts 25 Neocallimastigomycota Zygomycota ChytridiomycotaGlomeromycotaBasidiomycota Ascomycota Blastocladiomycota Microsporidia Fungi Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display.

26. Named for basidium – club-shaped sexual reproductive structure Karyogamy occurs within basidia Only diploid cell in life cycle Meiosis follows The four haploid products are incorporated into basidiospores 26

27. 27 Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. a. b. 2n n – Mating strain n + n Basidiocarp Basidium SterigmaBasidiospores MEIOSIS Zygote Primary mycelium (monokaryotic) Secondary mycelium (dikaryotic) + Mating strain KARYOGAMY Gills lined with basidia a: © Alexandra Lowry/The National Audubon Society Collection/Photo Researchers, Inc. FERTILIZATION

28. Spore germination leads to the production of monokaryotic hyphae Results in a monokaryotic mycelium, or primary mycelium Different mating types of monokaryotic hyphae may fuse Results in a dikaryotic mycelium, or secondary mycelium Heterokaryotic mycelium Basidiocarps (mushrooms) are formed entirely of secondary mycelium 28

29. Ascomycota Contain about 75% of the known fungi Includes bread yeasts, common molds, cup fungi, truffles, and morels Serious plant pathogens – cause of chestnut blight and Dutch elm disease Penicillin-producing fungi are in the genus Penicillium 29 Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Neocallimastigomycota Zygomycota ChytridiomycotaGlomeromycota Basidiomycota Ascomycota Blastocladiomycota Microsporidia Fungi

30. Named for ascus – microscopic, saclike reproductive structure Karyogamy occurs within asci Only diploid nucleus of life cycle Asci differentiate in ascocarp Meiosis and mitosis follow, producing 8 haploid nuclei that become walled ascospores 30

31. 31 Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. b. a. Developing mycelium Asexual reproduction Conidia Ascospore Each haploid nucleus divides once by mitosis Dikaryotic hyphae form From ascogonium Fully developed ascocarp composed of dikaryotic (ascogenic) hyphae and sterile hyphae (formation of young ascus) n + n + Mating strain Young ascus n 2n Ascogonium Conidia – Mating strain Antheridium c. MEIOSIS FERTILIZATION a: © Richard Kolar/Animals Animals; b: © Ed Reschke/Getty Images GERMINATION MITOSIS KARYOGAMY

32. Asexual reproduction is very common Conidia formed at the ends of modified hyphae called conidiophores Allow for the rapid colonization of a new food source Many conidia are multinucleate 32

33. Yeast Unicellular ascomycetes Most reproduce asexually by budding Yeasts can ferment carbohydrates Break down glucose into ethanol and CO 2 Used to make bread, beer, and wine Saccharomyces cerevisiae 33 Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. © David Scharf/Photo Researchers, Inc. 5 µm

34. Yeast is a long-standing model system for genetic research First eukaryotes to be manipulated extensively Saccharomyces cerevisiae first eukaryote to have genome sequenced Yeast two-hybrid system has been an important component of research on protein interactions 34

35. Ecology of Fungi Fungi, together with bacteria, are the principal decomposers in the biosphere Break down cellulose and lignin from wood Release carbon, nitrogen, and phosphorus 35

36. Fungi symbioses Obligate symbiosis – essential for fungus survival Facultative symbiosis – nonessential Interactions Pathogen – pathogens harm host by causing disease Parasites cause harm to host (do not cause disease) Commensal relationships benefit one partner but does not harm the other Mutualistic relationships benefit both partners 36

37. Endophytic fungi Live in the intercellular spaces inside plants Some parasitic, some commensalistic Some fungi protect their hosts from herbivores by producing toxins 37

38. Italian rye grass is more resistant to aphid feeding in the presence of endophytes 38 Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Fungal endophyteNo endophyte Fungal EndophyteNo Endophyte Aphids after 36 days 0 20 40 60 80 100 120 140 5 aphids SCIENTIFIC THINKING Hypothesis: Endophytic fungi can protect their host from herbivory. Prediction: There will be fewer aphids (Rhopalosiphum padi, an herbivore) on perennial ryegrass (Lolium perenne) infected with endophytic fungi than on uninfected ryegrass. Test: Place five adult aphids on each pot of 2-week-old grass plants with and without endophytic fungi. Place pots in perforated bags and grow for 36 days. Count the number of aphids in each pot. Result: Significantly more aphids were found on the uninfected grass plants. Conclusion: Endophytic fungi protect host plants from herbivory. Further Experiments: How do you think the fungi protect the plants from herbivory? If they secrete chemical toxins, could you use this basic experimental design to test specific fungal compounds? (left): © Nigel Cattlin/Alamy; (right): © B. Borrell Casal/Frank Lane Picture Agency/Corbis

39. Lichens Symbiotic associations between a fungus and a photosynthetic partner Cyanobacteria, green algae, or sometimes both Most are mutualistic Ascomycetes are found in all but about 20 of the 15,000 lichen species 39 Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Algal cells Fungal hyphae 40 μm © Ed Reschke

40. Fungi in lichens are unable to grow normally without their photosynthetic partners Fungi protect their partners from strong light and desiccation Lichens have invaded the harshest habitats Striking colors play a role in protecting photosynthetic partner Sensitive to pollutants 40 Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. a.b.c. Fruticose LichenCrustose LichenFoliose Lichen a: © Ken Wagner/Phototake; b: © Robert & Jean Pollock/Visuals Unlimited; c: © Robert Lee/Photo Researchers, Inc.

41. Mycorrhizae Mutualistic relationships between fungi and plants Found on the roots of about 90% of all known vascular plant species Function as extensions of root system Increase soil contact and absorption Two principal types Arbuscular mycorrhizae Ectomycorrhizae 41

42. Arbuscular mycorrhizae By far the most common Fungal partners are glomeromycetes No aboveground fruiting structures Potentially capable of increasing crop yields with lower phosphate and energy inputs 42 Hyphae penetrate the root cell wall but not plant membranes Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. © Eye of Science/Photo Researchers, Inc. a. Root 5 µm Arbuscular Mycorrhizae

43. Ectomycorrhizae Most hosts are forest trees (pines, oaks) Fungal partners are mostly basidiomycetes At least 5000 species of fungi are involved in ectomycorrhizal relationships 43 Hyphae surround but do not penetrate the root cells Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. © Dr. Gerald Van Dyke/Visuals Unlimited b. 50 µm Ectomycorrhizae

44. Animal mutual symbioses Ruminant animals host neocallimastigamycete fungi in their gut Leaf-cutter ants have domesticated fungi which they keep in underground gardens Ants provide fungi with leaves Fungi are food for the ants 44

45. Fungal Parasites and Pathogens Fungal species cause many diseases in plants Among most harmful pests of living plants Can also spoil harvested or stored food products 45 c. Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. b: © agefotostock/SuperStock; c: USDA Forest Service Archive, USDA Forest Service, Bugwood.org b.

46. 46 a. Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Courtesy of Ralph Williams/USDA Forest Service

47. Fungi may secrete substances making food unpalatable, carcinogenic, or poisonous Fusarium – vomitoxin Aspergillus flavus – aflatoxin 47 Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. a.b. 5 μm a: © Dayton Wild/Visuals Unlimited; b: © KAGE-Mikrofotografie

48. Fungi also cause human and animal diseases Candida – thrush; vaginal infections Pneumocystis jiroveci – pneumonia in AIDS Athlete’s foot, ringworm, and nail fungus Fungal diseases are difficult to treat because of the close phylogenetic relationship between fungi and animals 48

49. Batrachochytrium dendrobatidis causes chytridiomycosis 49 – Responsible for the worldwide decline in amphibian populations Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. Chytrid 10 μm (left): © School of Biological Sciences, University of Canterbury, New Zealand; (inset): Courtesy of Dr. Peter Daszak